Today is a special day for palaeognath publications, principally pertaining to the plethora of published PeerJ papers (well, three of them anyway) released today, featuring my team’s research! An early Crimbo comes this year in the form of three related studies of hind limb anatomy, development, evolution and biomechanics in those flightless feathered freaks of evolutionary whimsy, the ratites! And since the papers are all published online in PeerJ (gold open access), they are free for anyone with internet access to download and use with due credit. These papers include some stunning images of morphology and histology, evolutionary diagrams, and a special treat to be revealed below. Here I’ll summarize the papers we have written together (with thanks to Leverhulme Trust funding!):

My final year PhD student and “emu whisperer” Luis Lamas has published his first paper with co-supervisor Russ Main and I. Our paper beautifully illustrates the gross anatomy of the leg muscles of emus, and then uses exhaustive measurements (about 6524 of them, all done manually!) of muscle architecture (masses, lengths, etc.) to show how each of the 34 muscles and their tendons grew across a more than tenfold range of body mass (from 6 weeks to 18 months of age). We learned that these muscles get relatively, not just absolutely, larger as emus grow, and their force-generating ability increases almost as strongly, whereas their tendons tend to grow less quickly. As a result, baby emus have only about 22% of their body mass as leg muscles, vs. about 30% in adults. However, baby emus still are extremely athletic, more so than adults and perhaps even “overbuilt” in some ways.

This pattern of rapidly growing, enlarged leg muscles seems to be a general, ancestral pattern for living bird species, reflecting the precocial (more independent, less nest-bound), cursorial (long-legged, running-adapted) natural history and anatomy, considering other studies of ostriches, rheas, chickens and other species close to the root of the avian family tree. But because emus, like other ratites, invest more of their body mass into leg muscles, they can carry out this precocial growth strategy to a greater extreme than flying birds, trading flight prowess away for enhanced running ability. This paper adds another important dataset to the oft-neglected area of “ontogenetic scaling” of the musculoskeletal system, or how the locomotor apparatus adapts to size-/age-related functional/developmental demands as it grows. Luis did a huge amount of work for this paper, leading arduous dissections and analysis of a complex dataset.

Superficial layer of leg muscles in an emu, in right side view. Click any image here to emu-biggen. The ILPO and IC are like human rectus femoris (“quads”); ILFB like our biceps femoris (“hams”); FL, GM and GL much like our fibularis longus and gastrocnemius (calf) muscles, but much much bigger! Or, perhaps FL stands for fa la la la la?

Data for an extra set of emus studied by coauthor Russ Main in the USA, which grew their muscles similarly to our UK group. The exponents (y-axis) show how much more strongly the muscles grew than isometry (maintaining the same relative size), which is the dotted line at 1. The numbers above each data point are the # of individuals measured. Muscle names are partly above; the rest are in the paper. If you want to know them, we might have been separated at birth!

My second year PhD student Sophie Regnault (guest-blogger here before with her rhino feet post) has released her first PhD paper, on the evolution of kneecaps (patellae) in birds, with a focus on the strangeness of the region that should contain the patella in emus. This is a great new collaboration combining her expertise in all aspects of the research with coauthor Prof. Andy Pitsillides‘s on tissue histology and mine on evolution and morphology. This work stems from my own research fellowship on the evolution of the patella in birds, but Sophie has taken it in a bold new direction. First, we realized that emus don’t have a patella– they just keep that region of the knee extensor (~human quadriceps muscle) tendon as a fatty, fibrous tissue throughout growth, showing no signs of forming a bony patella like other birds do. This still blows my mind! Why they do this, we can only speculate meekly about so far. Then, we surveyed other ratites and related birds to see just how unusual the condition in emus was. We discovered, by mapping the form of the patella across an avian family tree, that this fatty tendon seems to be a thing that some ratites (emus, cassowaries and probably the extinct giant moas) do, whereas ostriches go the opposite direction and develop a giant double-boned kneecap in each knee (see below), whereas some other relatives like tinamous and kiwis develop a more “normal”, simple flake-like bit of bone, which is likely the state that the most recent common ancestor of all living birds had.

There’s a lot in this paper for anatomists, biomechanists, palaeontologists, ornithologists, evo-devo folks and more… plenty of food for thought. The paper hearkens back to my 2002 study of the evolution of leg tendons in tetrapods on the lineage that led to birds. In that study I sort of punted on the question of how a patella evolved in birds, because I didn’t quite understand that wonderful little sesamoid bone. And now, 12 years later, we do understand it, at least within the deepest branches of living birds. What happened further up the tree, in later branches, remains a big open subject. It’s clear there were some remarkable changes, such as enormous patellae in diving birds (which the Cretaceous Hesperornis did to an extreme) or losses in other birds (e.g., by some accounts, puffins… I am skeptical)– but curiously, patellae that are not lost in some other birds that you might expect (e.g., the very non-leggy hummingbirds).

Fatty knee extensor tendon of an emu, showing the absence of a patella. The fatty tissue is split into superficial (Sup) and deep regions, with a pad corresponding to the fat pad in other birds continuous with it and the knee joint meniscus (cushioning pad). The triceps femoris (knee extensor) muscle group inserts right into the fatty tendon, continuing on over it. A is a schematic; B is a dissection.

Sectioning of a Southern Cassowary’s knee extensor tendon, showing: A, Similar section as in the emu image above. revealing similar regions and fibrous tissue (arrow), with no patella, just fat; and B, With collagen fibre bundles (col), fat cells (a), and cartilage-like tissue (open arrows) labelled.

Evolution of patellar form in birds. White branches indicate no patella, blue is a small flake of bone for a patella, green is something bigger, yellow is a double-patella in ostriches, black is a gigantic spar of bone in extinct Hesperornis and relatives, and grey is uncertain. Note the uncertainty and convergent evolution of the patella in ratite birds (Struthio down to Apteryx), which is remarkable but fits well with their likely convergent evolution of flightlessness and running adaptations.

Finally, Kyle Chadwick came from the USA to do a technician post and also part-time Masters degree with me on our sesamoid grant, and proved himself so apt at research that he published a paper just ~3 months into that work! Vivian Allen (now a postdoc on our sesamoid bone grant) joined us in this work, along with Sophie Regnault. We conceived of this paper as fulfilling a need to explain how the major tissues of the knee joint in ostriches, which surround the double-patella noted above, all relate to each other and especially to the patellae. We CT and MRI scanned several ostrich knees and Kyle made a 3D model of a representative subject’s anatomy, which agrees well with the scattered reports of ostrich knee/patellar morphology in the literature but clarifies the complex relationships of all the key organs for the first time.

This ostrich knee model also takes Kyle on an important first step in his Masters research, which is analyzing how this morphology would interact with the potential loads on the patellae. Sesamoid bones like the patella are famously responsive to mechanical loads, so by studying this interaction in ostrich knees, along with other studies of various species with and without patellae, we hope to use to understand why some species evolved patellae (some birds, mammals and lizards; multiple times) and why some never did (most other species, including amphibians, turtles, crocodiles and dinosaurs). And, excitingly for those of you paying attention, this paper includes links to STL format 3D graphics so you can print your own ostrich knees, and a 3D pdf so you can interactively inspect the anatomy yourself!

Ostrich knee in side view: A, X-ray, and (B) labelled schematic.

3D model of an ostrich knee, showing: A, View looking down onto the top of the tibia (shank), with the major collateral ligaments (CL), and B, View looking straight at the front of the knee joint, with major organs of interest near the patella, sans muscles.

You can view all the peer review history of the papers if you want, and that prompts me to comment that, as usual at PeerJ (full disclosure: I’m an associate editor but that brings me £0 conflict of interest), the peer review quality was as rigorous at a typical specialist journal, and faster reviewing+editing+production than any other journal I’ve experienced. Publishing there truly is fun!

Merry Christmas and Happy Holidays — and good Ratite-tidings to all!

And stay tuned- the New Year will bring at least three more papers from us on this subject of ratite locomotion and musculoskeletal anatomy!

♬Should auld palaeognathans be forgot, And never brought for scans? Should publications be soon sought, For auld ratite fans!♪

It’s World Penguin Day! Watch your back though… these penguins aren’t as nice as they seem. But they need us to be nice to them!

Whether you watch a classic GIF like the one above, or a kid-friendly TV/film documentary, you might get the impression that penguins lead carefree, or at least silly or slapstick, lives– happy feet and all that. It works for Hollywood: a Charlie Chaplin comedy relief role to play. And that’s the vision of penguins I grew up with: they were living cartoons to me.

But what’s the reality? Plenty of documentaries, most notably to my mind the recent Attenborough’s “Frozen Earth” episodes or “March of the Penguins” film, have dealt with the darker side to these two-toned, tuxedo-toting antipodeans. And anyone who has experienced penguins in the wild has probably seen those not-so-light facets of penguinity firsthand. On realiizing just how compulsively horny young “hooligan cock” male penguins were, Natural History Museum ornithologist Douglas Russell wrote: ““just the frozen head of the penguin, with self-adhesive white O’s for eye rings, propped upright on wire with a large rock for a body, was sufficient stimulus for males to copulate and deposit sperm on the rock.”

Stomach-Churning Rating: 5/10; some tears may be shed over cute baby penguins and you might choke if you’re a rhea trying to swallow one, but the anatomy shown is mostly skeletal or dessicated. No penguin juices. Except those just mentioned above.

I’m quick to admit, I didn’t know much about penguins until recently. I couldn’t name many species or say much about their behaviour, anatomy or evolutionary history. When I was a graduate student at Berkeley, I was enthused by a now-classic, elegantly simple study (published in 2000) that fellow PhD student Tim Griffin and biomechanist Dr. Rodger Kram conducted on penguin waddling. They found that the waddling gait of penguins isn’t mechanically disadvantageous, as it appears, but rather is a way that they conserve energy while walking. It’s the short legs, instead, that make their gait metabolically expensive, because shorter legs mean that more frequent, costly steps need to be taken, incurring high costs due to rapid firing of leg muscles to support the body. My vicarious enjoyment of Griffin’s & Kram’s research began my scientific introduction to penguins. Fast forward to 2014: I get a crash course in penguinology.

Mostly-fledged Magellanic penguin

That’s what this post is about, and how it brought me in touch with The Existentialist Penguin– the haggard, storm-tossed, predator-harried, starved and bullied wanderer of wastelands.

My personal introduction to penguins over the past year has been initiated by a collaboration with PhD student James Proffitt and long-time colleague Dr. Julia Clarke, both at the University of Texas in Austin. They kindly invited me to collaborate on applying modern biomechanics to the surprisingly excellent fossil record of penguins (Sphenisciformes), among other extant water birds. Before diving into it all, I happened to go to Argentina.

Penguin tries to keep cool in the shade, opening its mouth to shed heat in the autumn sun.

Just before I travelled to Patagonia on unrelated business (to study sauropodomorph dinosaurs!), I did a little googling and came across Punta Tombo reserve, near the city of Trelew that I was visiting (more about that in a future post!). It’s where some 1+ million Magellanic penguins (Spheniscus magellanicus) gather every southern summer to breed and fledge before making a long ~5 month swim up to Brazil. I asked my host, Dr. Alejandro Otero, if we might take a day off to visit this spot, where guanacos, rheas and other wildlife were also said to be common, and he basically said “Hell yes!” as he’d never been there. My Flickr photostream gives a big set of my favourite photos from that trip, but here are some others below, to show some of my experiences. We rented a car and took a lovely 90-minute drive south across the Patagonian plains, observing wildlife like tinamous (yes! So exciting for me) as we went. You could get within 1.5m of the penguins according to park rules, and the penguins were very permissive of that!

This jaunty chap was staying put in his burrow while people walked by. We came closer and he kept rotating his head around, staring at us. I first took it as cute juvenile behaviour, but on later observations of penguins realized it was a threat- “My beak is sharp! Stay back, bro, or I’ll glock ya!”

The video below shows a penguin encounter that left me with no doubts that these animals don’t mess around. The smaller penguin escaped, losing its cool burrow and some of its tough hide, too. Indeed, penguins can be remarkable assholes to each other.

With battles like this erupting all around us, where the penguins struggled to find shade in the desert-like inland parts of the park, often hundreds of meters away from the cool ocean, it came as no surprise to find casualties. The juveniles (and some remaining adults; most having left by now while the ~1 year-old juveniles fledge) not only battled, but also fasted, and roasted in the heat as they shed their insulatory fluff for waterproofed streamlining. This poor little flat Spheniscus had been trodden a bit past streamlined:

Near the end of our visit, just after I saw an informative sign about the lesser rhea or “choique” (Pterocnemia/Rhea pennata), we managed to get very close to a rhea and follow it for a while, as penguins stood around in apparent disinterest. I’ll never forget that meeting: two flightless birds, yet adapted to such different lifestyles and habitats. The penguins were in the rhea’s domain; a hot, wind-blown, scree-scoured scrubland on the edge of the fertile ocean.

The choique soon found a dry old hatchling penguin carcass, no meatier than the surrounding thickets, and tried to swallow it. The loss of teeth by its distant ornithurine ancestors proved to be a bad move, because it struggled to get the jerky-like mass through its beak:

That Punta Tombo visit was an experience I’ll never forget. I returned to the UK, abuzz with excitement about penguins. I “got” them now, I felt, at least in a very unscientific, anthropomorphic way. It took the face-to-beak experience to drive that home, more than any emotive film treatment could. Whether enduring Antarctic wintery blasts or unforgivingly hot and dry, burrow-speckled coastal badlands, penguins are buggers with true grit. Survivors, as their >60 million year fossil record attests to. On my return, I delved through my photos of museum specimens to get a better appreciation for penguin anatomy, preparing to also get familiar with that fossil record; all as part of that ongoing work with Proffitt and Clarke. Here’s some of that anatomy:

My first encounter with a penguin in the wild (but not a live one) is probably this specimen washed up on a beach in Uruguay. I’m going with the tentative ID of a juvenile penguin skeleton (short foot; flat wing bones); probably Magellanic. The bevy of vertebrate morphologists investigating dead penguins on this beach during our conference in 2010 will not soon be forgotten!

University Museum of Zoology Cambridge skeleton of a “great penguin” (do not confuse with the great pumpkin!) species of Aptenodytes; either King (patagonicus) or Emperor (forsteri). Characteristic features, in addition to the robust, dense skeleton, include the short neck, flattened but robust wings and scapulae, robust furcula (wishbone), stubby legs (with a big blocky patella) and thin but longish tail (supposedly used to balance with while walking/standing).

I’ll visit some more penguin anatomy in coming images- those photos are just teasers. And they set the stage for me to go back to my one-stop-shopping for awesome ornithological specimens, the Natural History Museum at Tring (images below presented with kind permission from the Natural History Museum, London; but I took the photos), to pick up an assortment of 11 frozen penguins from helpful curator Hein van Grouw! Such as this “gagged” King penguin:

And this handsome Emperor penguin, going through the Equine Imaging Centre’s CT scanner as I do my usual routine of (1) get cool critters, (2) barrage them with radiation to peek inside:

CT scanner monitors as I scan a penguin; mid-torso x-ray slice shown on the right.

Because I love the CT scan images of these penguins so much (their skeletons are awesome and bizarre!), I’ll share the pilot scans of the best ones now:

Calling all penguin experts! What’s up with this? Is that really how much gastrolith (stomach stone; near bottom of image) volume a penguin carries (answer after some literature reading: maybe yes!), or did a museum curator stick rocks up its bum? It seems very caudal in position, and this is consistent with other animals I’ve seen (some below). A paper on this phenomenon and potential role in ballast is here. Another here.

Side view. Nice view of the head at least.

Young juvenile. Nice pose, and lots of anatomy is shown. And check it out- gastroliths?!? In such a young animal– is it even feeding yet? (presumably straight after hatching) And they are relatively big pebbles, too! If I noticed this 5 years ago, it would have been a nice paper to report- first recognition of gastroliths in penguin chicks seems to have been then. Indeed, that study observed some chicks intentionally swallowing stones.

Another youngun; the fluffy one from the photo above. More rocks up its wazoo.

Three wee little chicks, all with stomach stones.

CT reconstruction of adult skeleton. This specimen was gutted and flattened, so the gastroliths are few and scattered. Check out the long tail:

From recent skeletons to fossil ones, penguins have wacky anatomy; they break most of the “rules” of being a proper bird, putting other oddballs like rheas to shame. I can’t ably review the many penguin species we know of, but the ancient Palaeocene penguin Waimanu features prominently in recent scientific discussions of penguin evolution, such as the superb research and blog of Dan Ksepka as well as many workers in the southern hemisphere. I haven’t had a chance to inspect that creature’s bones, but while in Trelew, Argentina, I was very pleased to run into some excellent specimens of a later animal:

Part of the nice skeleton of Palaeospheniscus patagonicus, an Oligocene/Miocene largish penguin; from the MEF collections in Trelew, Argentina and collected nearby. The genus has been known since Ameghino’s description in 1891, and is closely related to living penguins, especially Aptenodytes. It was not a large penguin, but at about 5kg body mass was no slouch as birds go (roughly similar in size to a Magellanic penguin). I also got to see Madrynornis mirandus, a Miocene form.

For me, the diagnostic trait of a penguin skeleton: the very short, tobust tarsometatarsus. From Palaeospheniscus, as above. The great palaeontologist GG Simpson wrote of it: “Despite the innumerable variations in details, the tarsometatarsi, on which all species but P. robustus are based, are quite stereotyped in general structure and leave little doubt that the forms placed here by Ameghino do all belong to a natural group.” A ratio of length to proximal width of >2 is typical of most penguins. Synapomorphy FTW!

From beach skeletons, to mass suffering of landbound birds, to 3D imaging and fossil skeletons, I’ve had quite the immersion in penguinness lately. And through that experience, I’ve been drawn closer to penguins in more ways than one. I’ve been impressed by their adaptability and durability. In some ways, penguins’ adaptations to harsh freezing winters in wastelands also aid them to survive harsh baking summers in dry badlands.

Yes, those badlands are still coastal, and penguins can still drink the saltwater and excrete salt via their supraorbital glands, but those penguins in Punta Tombo were not having a keg party. They were clearly enduring some serious discomfort, and not all making it through the ordeal. I watched silently along with other penguins as one penguin lay prone in an awkward pose on a bleached-white stretch of hardpan soil, while one flipper meekly raised, then flopped down. It was not long for this world, and there was a host of large scavengers around ready to make the most of that, while penguin-eating giant petrels (a sister group to penguins) wheeled overhead.

Waddlers of the wastes

While penguins still spend most of their lives at sea, they retain a sometimes astonishing array of behaviours they use on land: burrowing, hopping/jumping, costly short-legged (but efficiently waddling) walking, and perhaps more that we haven’t yet discovered! Their unique anatomy reflects a compromise between all these factors, and we’re fortunate to have knowledge of their fossil record that shows a lot of detail on how they evolved it all. While penguins are a highly aquatic species, they show how aquatic and terrestrial adaptations can coexist in harmony; it’s not just a black-or-white issue. But with climate change in progress, the ~18 species of penguins have some rapidly altering challenges to adapt to, or go the way of Waimanu. This is a critical Kierkegaardian moment for The Existentialist Penguin.

I raise a glass in toast to that versatile, resilient, gravel-gizzarded Existentialist Penguin! May it persevere all the troubles our ever-changing world throws at it, as it has done since the Palaeocene. And may we draw inspiration from its tenacity, to face our own troubles, together on this crazy spinning globe!

(John: here’s a guest post from my former PhD student, soon to be 100% legit PhD, Dr., and all that jazz, Julia Molnar!)

This is my first guest post, but I have been avidly following what’s in John’s freezer (and the blog too) for quite a while. I joined the lab in 2009 and left a month ago on the bittersweet occasion of surviving my PhD viva (oral exam/defense), so I’d like to take a moment here to thank John and the Structure & Motion Lab for a great 4 years!

Moving on to freezer-related matters; specifically, a bunch of frozen crocodile spines. It was late 2011, and the reason for the spines in John’s freezer was that John, Stephanie Pierce, and I were trying to find out more about crocodile locomotion. This was anticipated to become my first major, first-author research publication (but see my Palaeontologia Electronica paper on a related subject), and I was about to find out that these things seldom go as planned; for example, the article would not be published for more than three years (the research took a long time!). Before telling the story of how it lurched and stumbled toward eventual publication, I’ll give you some background on the project.

First of all, why crocodiles? For one thing, they’re large, semi-terrestrial animals, but they use more sprawling postures than typical mammals. Along with alligators and gharials, they are the only living representatives of Crocodylomorpha, a 200+ million year-old lineage that includes wolf-like terrestrial carnivores, fish-like giants with flippers and a tail fin, even armored armadillo-like burrowers. Finally, crocodiles are interesting in their own right because they use a wide variety of gaits, including bounding and galloping, which are otherwise known only in mammals.

Nile crocodile skeletal anatomy

OK, so why spines? Understanding how the vertebral column works is crucial to understanding locomotion and body support on land, and inter-vertebral joint stiffness (how much the joints of the backbone resist forces that would move them in certain directions) in particular has been linked to trunk movements in other animals. For this reason, vertebral morphology is often used to infer functional information about extinct animals, including dinosaurs. However, vertebral form-function relationships have seldom been experimentally tested, and tests on non-mammals are particularly scarce. So we thought the crocodile spines might be able to tell us more about the relationship between vertebral morphology, mechanics, and locomotion in a broader sample of vertebrate animals. If crocodile spine morphology could be used to predict joint stiffness, then morphological measurements of extinct crocodile relatives would have some more empirical heft to them. Several skeletal features seem to play roles such as levers to mechanically stiffen crocodile spines (click to emcroc’en):

Anatomy of a crocodile vertebra

We decided to use a very simple technique that could be replicated in any lab to measure passive stiffness in crocodile cadavers. We dissected out individual joints were and loaded with known weights. From the movement of the vertebrae and the distance from the joint, we calculated how much force takes to move the joint a certain number of degrees (i.e. stiffness).

Me with crocodile vertebra and G-clamp

X-ray of two crocodile vertebrae loaded with a metric weight to calculate their joint’s stiffness

Afterwards, we boiled the joints to remove the soft tissues – the smell was indescribable! We took 14 measurements from each vertebra. All of these measurements had been associated with stiffness or range of motion in other studies, so we thought they might be correlated with stiffness in crocodiles also.

Some of the vertebral measurements that were related to stiffness

Despite my efforts to keep it simple, the process of data collection and analysis was anything but. I recall and exchange with Stephanie Pierce that went something like this:

Stephanie: “How’s it going?”

Me: “Well, the data are messy, I’m not seeing the trends I expected, and everything’s taking twice as long as it was supposed to.”

Stephanie: “Yes, that sounds like science.”

That was the biggest lesson for me: going into the project, I had been unprepared for the amount of bumbling around and re-thinking of methods when the results were coming up implausible or surprising. In this case there were a couple of cool surprises: for one thing, crocodiles turn out to have a very different pattern of inter-vertebral joint stiffness than typical mammals: while mammals have stiff thoracic joints and mobile lumbar joints, crocodiles have stiffer lumbar joints. Many mammals use large lumbar movements during bounding and galloping, so crocodiles must use different axial mechanics than mammals, even during similar gaits. While that’s not shocking (they did evolve their galloping and bounding gaits, and associated anatomy, totally independently), it is neat that this result came out so clearly. Another unexpected result was that, although several of our vertebral measurements were correlated with stiffness, some of the best predictors of stiffness in mammals from previous studies were not correlated with stiffness in crocodiles. The study tells a cautionary tale about making assumptions about extinct animals using data from only a subset of their living relatives or intuitive ideas about form and function.

Finally, the experience of doing the experiments and writing the paper got me interested in other aspects of crocodilian functional anatomy. For instance, how does joint stiffness interact with other factors, such as muscle activity and properties of the ribs, skin, and armor in living crocodiles? Previous studies by Frey and Salisbury had commented on this, but the influence of those factors is less tractable to experiment on or model than just naked backbones with passively stiff joints. In the future, I’d like to study vertebral movements during locomotion in crocodiles – especially during bounding and galloping – to find out how these patterns of stiffness relate to movement. In the meantime, our study shows that, to a degree, crocodile backbone dimensions do give some clues about joint stiffness and locomotor function.

To find out more, read the paper! It was just featured in Inside JEB.

Julia Molnar, Stephanie Pierce, John Hutchinson (2014). An experimental and morphometric test of the relationship between vertebral morphology and joint stiffness in Nile crocodiles (Crocodylus niloticus). The Journal of Experimental Biology 217, 757-768 link here and journal’s “Inside JEB” story

And I post my blog and stareInto x-rays of an ostrichI’ve always known that radiographs never liePeople always say “that’s cool”To see x-rays of an ostrichSo keen to know whatLies behind the skin

(evolved from “Eyes of A Stranger” by Queensrÿche, from the epic masterpiece of Operation: Mindcrime (1988). One of my favourite albums of all time, and a fantastic concept album too. The band was operating at their peak. Tight! Drug addict Nikki gets brainwashed by the evil Dr. X and made to assassinate a nun, Sister Mary, who was a prostitute, and then there’s like a revolution or something, and things get all screwed up and no one ends up happy – or alive. All the while, Geoff Tate is singing his guts out. Anyway, I got to see them play the whole album live in 1990 in Madison, WI, for the filming of Operation: Livecrime, which was like a Mecca moment for me back then. Look for me (pre-bald years) in about the 6th row. )

What does that album have to do with the number 2 (two days left in Freezermas)? Hmm… Track 2 is the instrumental Anarchy-X, and today’s post is about X-rays as well as that funky ostrich (2 legs good! 2 toes good, too!) again, so I’m satisfied, and by this point you’re probably just oggling the mind-blowing images below anyway, so fuck it!

Stomach-Churning Rating: 2/10; just X-rays.

Tech/MRes Kyle Chadwick, Renate Weller and the equine imaging team at the RVC took these x-rays of our birdie for us and for an artist who is doing a big x-ray animal art show (more news on this soon!)– thanks to all of them for some truly awesome images! I could stare at the intricate details in these images for hours– go ahead, do it. Click to emostrichinate them (this post needs to be viewed on nice big screen), and oggle away…

Head and neck.

Another view of the same. The highly flexible esophagus and trachea can be seen going diagonally across the neck; twisting from ventral to dorsal. It’s floppy, so it can do that.

Neck near the head; tapering.

Middle of neck. Check out the rings of the trachea!

Base of neck and shoulder.

Shoulder and chest. Hard to image; thick and dense (still was frozen), hence the whiteout toward the left side of the image.

Check out that wing!!

Ankle- note the big calloused pad that ostriches rest on (right side of image).

That two-toed foot… but did you know that normally the missing 2nd toe is still there as a fibrous remnant on the 3rd toe?

Freezermas continues! Today we have a treat for you. Lots of detailed anatomy! This post comes from my team’s dissections of an ostrich last week (~3-7 February 2014), which I’ve been tweeting about as part of a larger project called the Open Ostrich.

However, before I go further, it’s as important as ever to note this:

Stomach-Churning Rating: 9/10: bloody pictures of a dissection of a large ostrich follow. Head to toes, it gets messy. Just be glad it wasn’t rotten; I was glad. Not Safe For Lunch!

If the introductory picture below gets the butterflies a-fluttering in your tummy, turn back now! It gets messier. There are tamer pics in my earlier Naked Ostriches post (still, a rating of 6/10 or so for stomach-churning-ness there).

Research Fellow Jeff Rankin, myself and technician/MRes student Kyle Chadwick get to work, removing a wing.

This is a male ostrich, 71.3 kg in body mass, that had gone lame in one foot last summer and, for welfare reasons, we had to put down for a local farmer, then we got the body to study. We took advantage of a bad situation; the animal was better off being humanely put down.

The number for today is 6; six posts left in Freezermas. But I had no idea I’d have a hard time finding a song involving 6, from a concept album. Yet 6 three times over is Slayer’s numerus operandi, and so… The concept album for today is Slayer’s 1986 thematic opus “Reign in Blood” (a pivotal album for speed/death metal). The most appropriate track here is the plodding, pounding, brooding, then savagely furious “Postmortem“, which leads (literally and figuratively, in thunderous fashion) to the madness of the title track, after Tom Araya barks the final verse:

“The waves of blood are rushing near, pounding at the walls of lies

Turning off my sanity, reaching back into my mind

Non-rising body from the grave showing new reality

What I am, what I want, I’m only after death”

I’m not going to try to reword those morbid lyrics into something humorous and fitting the ostrich theme of this post. I’ll stick with a serious tone for now. I like to take these opportunities to provoke thought about the duality of a situation like this. It’s grim stuff; dark and bloody and saturated with our own inner fears of mortality and our disgust at what normally is politely concealed behind the integumentary system’s viscoelastic walls of keratin and collagen.

But it’s also profoundly beautiful stuff– anatomy, even in a gory state like this, has a mesmerizing impact: how intricately the varied parts fit together with each other and with their roles in their environment, or even the richness of hues and multifarous patterns that pervade the dissected form, or the surprising variations within an individual that tell you stories about its life, health or growth. Every dissection is a new journey for an anatomist.

OK I’ve given you enough time to gird yourself; into the Open Ostrich we go! The remainder is a photo-blog exploration of ostrich gross anatomy, from our detailed postmortem.

Here, I give you a long-planned post on the patella (“kneecap bone”) of birds, which was my Royal Society Senior Research Fellowship sabbatical project for 2012-13. This is only a brief introduction to the anatomical issues at hand, err, I mean at knee…

Stomach-Churning Rating: 6/10; mostly skeletons/fossils, but there are a few images of the dissection of a guineafowl, which is fresh and meaty.

Archaeopteryx, the Berlin specimen. Helluva fossil, but nary a patella!

The question I am exploring, first of all, is simply how the patella evolved, because it seems to be present in almost all living birds. However, it is absent in all non-avian dinosaurs, and indeed most Mesozoic birds, too. There is barely a hint of any precursor structure (a “patelloid”) in other reptiles, but lizards evolved their own patella that is quite different (a flattened lozenge, not a rectangular structure lying tightly confined in a “patellar groove” on the femur as it is in birds). Mammals evolved the knobbly, hemispherical kind of kneecap that we’re familiar with, possibly on several occasions (a different story!). So the patella evolved at least three times in the lizard, mammal and bird lineages– and possibly more than once in each of these groups. And that’s about it for almost 400 million years of tetrapod evolution, except for a few very rare instances in fossils and sort-of-patella-like things in some frogs or other weirdos.

Fossil birds exhibit no clear presence of a patella until we come very close to modern birds on the avian stem of the tree of life (see below). And then, suddenly in modern birds, there is a lot of variation and not much good documentation of what kind of patella exists. This makes it challenging to figure out if the patella is ancient for modern birds or if it evolved multiple times, or how it changed after it first evolved– let alone bigger questions of what the patella was “for” (performance benefits, functional consequences, etc.; and developmental constraints) in the birds that first evolved it.

Considering that the patella is such an obvious bone in some birds, and certainly affects the mechanics of the knee joint (forming a lever for the muscles that cross it; homologous to our quadriceps muscles) and hence locomotion, it is a compelling research topic for me.

What follows is a pictorial guide to the patella of some birds, in sort of an evolutionary/temporal sequence (see my earlier post for a recap of some major groups), with a focus on animals I’ve studied more intensively so far (with >10,000 species, there is a lot that could be done):

The early Cretaceous bird Gansus (from the IVPP in Beijing), represented by many beautifully preserved specimens, all of which lack a patella. This absence is characteristic of other stunningly preserved fossil Chinese birds, indicating that this is almost certainly an ancestral absence of a patella, until…

The famed Cretaceous diving (flightless) bird Hesperornis, from Wikipedia/Smithsonian. Note the massive, conical/crested patella in front of the knee (jutting up and overlapping the ribs/vertebrae close to the pelvis; see also below). That elongate patella is characteristic of many diving birds that use foot-propelled swimming; it has evolved many times in this fashion. Other hesperornithiform birds show some transformational states in their anatomy toward this extreme one.

Check this out! More Hesperornis (cast), with the femur on the left and the patella on the right. The bloody patella is almost as long as the femur! That’s nuts. With kind permission from the Natural History Museum, London.

Exhibited ostrich (Struthio camelus) skeleton in left side view showing the patella (white arrow), on exhibit atThe Natural History Museum at Tring, Hertfordshire, UK. Ostriches are remarkable in that they have this elongate patella (actually a double patella; there is a smaller, often-overlooked second piece of bone) and yet are rather basal (closer to the root of the modern avian family tree)– however, they obviously are specialized in ways other than this double patella, most notably their very large size, flightlessness, and elongate legs. So the unusual patella is more likely linked to their odd lifestyle than a truly primitive trait, at least to some degree (but stay tuned: what happened with the patella in other members of their lineage, the ratites/palaeognaths, is much less well understood!).

Note that ostriches and Hesperornis together hint that the presence of a patella might have been an ancestral trait for living birds, but their patellae are so different that the ancestral state from which they evolved must have been different, too; perhaps simpler and smaller. Hence we need to look at other birds…

Skinned right leg of a Helmeted Guineafowl, Numida meleagris, above. That whitish band of tissue in the middle of the screen, on the front of the knee, is part of what is concealing the patella. That is an aponeurosis (connective tissue sheet, like a thin tendon) of the muscles corresponding to our “quads” or our tensor fascia latae, detailed more below. Guineafowl are fairly basal and well-studied in terms of their bipedal locomotion, so they are an important reference point for avian form and function.

Right guineafowl leg, with patella exposed. Here I’ve peeled away that white band of tissue and associated muscles, which have been reflected toward the bottom of the screen (AIL and PIL labels corresponding to the anterior and posterior parts of the Iliotibialis lateralis muscle). The tip of the scalpel is contacting the patella. It’s not much to see, but lies atop the bright yellow fat pad that cushions it against the femur. You should be able to see a groove in the end of the femur just above that fat pad, which is where the patella sits and slides up and down as the knee moves/muscles contract. This is called the patellar groove, or sulcus patellaris.

Left leg of another guineafowl (with right tibiotarsus behind it, on the left) showing the patellae in articulation; in medial (inside) and cranial (front) views, respectively. The patella is the little rectangular bit of bone in the top middle of the screen, interposed between femur (thigh) and tibiotarsus (shank).With kind permission from the Natural History Museum, London.

Right leg of a Cape Penguin (Spheniscus demersus) from the University Museum of Zoology in Cambridge, showing the big lumpy patella in this wing-propelled diver. They still walk long distances on land, so presumably a patella plays some role in their gait, helping to explain its large size, which like the ostrich and Hesperornis seems to be a novel trait. Notice the groove across the patella, made by the tendon of the ambiens (like our sartorius/”tailor’s muscle”), which crosses from the inside to the outside of the leg via this route. This groove is often considered a useful phylogenetic character in modern birds, as its contact with the patella (sometimes via a hole, or foramen) varies a lot among species.

A hornbill, Buceros sp., from the UMZ Cambridge museum as well. This displays the possibly-more-typical, little rounded patellar nubbin that many birds have. See below for more.

Closeup of the knee/patella of the hornbill, Buceros sp., from above. Not much to squawk about, patella-wise, but it’s there.

And so we complete our quick tour of the avian patella, in its grand variation and humble beginnings.

Why does an ostrich have a patella and a Tyrannosaurus, Edmontosaurus or Triceratops did not? Why were birds the only bipedal lineage to evolve a patella (mammals and lizards gained a patella as small quadrupeds), and why did some bipeds like kangaroos “lose” (reduce to fibrous tissue, apparently) their patella?

These are the kinds of mysteries my group will now be tackling, thanks to a generous Leverhulme Trust grant on sesamoid bone ontogeny, mechanics and evolution. My group is now Dr. Vivian Allen as the postdoc, Sophie Regnault as the PhD student, and Kyle Chadwick as the technician and MRes student, along with numerous collaborators and spin-off projects. We’re looking forward to sharing more! But for now, I hope that I’ve engendered some appreciation for the avian patella, as the silly title indicates (“fella” used in the general sense of anyone!). This work is all unpublished, but some of this should be out in not too long, in much more lavish detail! Much as the patella is the “forgotten lever “of the avian hindlimb, it is the fulcrum about which a substantial part of my research group’s activity now pivots.

Yesterday I encountered the question that, as a scientist who has studied a certain chunky Cretaceous carnivore a lot, most deflates me and makes me want to go study cancer therapeutic methods or energy sources that are alternatives to fossil fuels (but I’d be useless at either). I will explain why this is at the end of the post.

The question stems from a new discovery, reported in Proceedings of the National Academy of Sciences (PNAS) and thus expected to be one of the more important or exciting studies this year (no, I’m not going to get into the issue here of whether these “high impact” journals include the best scientific research or the most superficial or hyped “tabloid” science; they publish both, and not in mutual exclusivity). It’s a broken Tyrannosaurus rex tooth embedded in a duckbill dinosaur’s tail bone, which healed after the injury, showing that the animal survived the attack.

If you’re with me so far, you might be making the logical leap that this fossil find is then linked to the hotbed of furious controversy that still leaves palaeontology in crisis almost 100 years after Lambe suggested it for the tyrannosaur Gorgosaurus. If the hadrosaur survived an attack from a T. rex, then T. rex was a habitual predator and OMG JACK HORNER AND OTHERS BEFORE HIM WERE WRONG!

And you’d be right.

My encounter with the question stemmed from an email from a science journalist (Matt Kaplan) that, as is normal practice, shared a copy of the unpublished paper and asked for comments from me to potentially use in an article he was writing for the science journal Nature’s news site. Here, then, was my off-the-cuff response:

“Ooh. I do have a pretty strong opinion on this. Not sure if you’d want to use it but here goes. I may regret it, but this hits my hot buttons for One of the Worst Questions in All of Palaeobiology!

The T. rex “predator vs. scavenger” so-called controversy has sadly distracted the public from vastly more important, real controversies in palaeontology since it was most strongly voiced by Dr Jack Horner in the 1990s. I find this very unfortunate. It is not like scientists sit around scratching their heads in befuddlement over the question, or debate it endlessly in scientific meetings. Virtually any palaeontologist who knows about the biology of extant meat-eaters and the fossil evidence of Late Cretaceous dinosaurs accepts that T. rex was both a predator and scavenger; it was a carnivore like virtually any other kind that has ever been known to exist.

While the discovery is nice evidence, it is not particularly exciting in a scientific sense and is only one isolated element from species that lived for hundreds of thousands of years, which to me changes nothing and allows no generalizations about the biology of any species, only the statement that at one point in time a Tyrannosaurus bit a hadrosaur that survived the encounter. There is no real substance to the controversy that T. rex was “either” a predator or scavenger. It is just something that scientists drum up now and then to get media attention. I hope that soon we can move on to more pressing questions about the biology of extinct animals, but the media needs to recognize that this is just hype and they are being played in a rather foolish way; likewise scientists that still feel this is an exciting question need to move on. Maybe this specimen will allow that. But somehow my cynical side leads me to suspect that this “controversy” will just persist because people want it to, regardless of logic or evidence. (bold font added; see below)

Great galloping lizards, I am so tired of this nonsense. Maybe there is educational value in showing how science deals with provocative half-baked ideas about celebrity species, but scientists in the community need to speak up and say what the real science is about. It’s not about this “controversy”. Modern palaeontology is so much better than this.

Sorry for the rant. Maybe it’s too extreme but I’m just fed up with this non-issue! I suspect a huge proportion of our field feels similarly, however.”

(I later redacted a bit of it where I got a little too excited and used the word “curmudgeon”; a mistake, as that could be seen as ad hominem rather than a term of endearment, and this issue is about the science and not the people, per se. That bit is redacted here, too. I’ve also redacted a sentence in which I made an opinion on whether the paper should have been published in PNAS; that is mostly irrelevant here. I was not a reviewer, and authors/reviewers/editors have to make that decision. This would be a massive tangent away from what this blog post is intended to be about! I know some of the authors and don’t want to offend them, but this is about the science and how it is represented to the world, not about these particular authors or even this paper itself.)

Importantly, Kaplan’s story did include my skeptical quote at the end. I am curious to see how many other news stories covering this paper go that far.

Would a T. rex prey on, or just scavenge — or have a great time racing — a giant chicken? (art by Luis Rey)

I will stop right here and acknowledge that I’ve published a lot on a somewhat related topic: how fast a T. rex could run or if it could run at all. To me, that’s a great scientific question that has consequences not only for the predator/scavenger false dichotomy, but also for general theories of locomotor biomechanics (can an animal the size of a large elephant run as well as or better than said elephant? What are the thresholds of size and maximal running/jumping/other athletic abilities and how do they vary in different evolutionary lineages? And so on.). I’ll defend the validity of that question to the bitter end, even if it’s a question I’ve grown a little (but only a little) tired of and generally feel is about as well settled as these things can be in palaeontology (see my review here). I’ll also defend that it has been a real controversy (I have plenty of old emails, formal rebuttals submitted by colleagues, and other discourse as evidence of this) since I tackled it starting in 2002 and sort of finishing by 2011. I am sensitive about the issue of hyping my research up– this is something I’ve been careful about. I set a reasonable bar of how much is too much, check myself continuously with reflective thought, and I do not feel I have ever really crossed that bar, away from science-promotion into darker realms. This is partly why I’ve stopped addressing this issue in my current work. I feel like the science we’ve done on this is enough for now, and to keep beating the same drum would be excessive, unless we discovered a surprising new way to address the questions better, or a very different and more compelling answer to them.

“T. rex: scavenger or predator?” was controversial back in 1994 when Horner published “The Complete T. rex”, where he laid out his arguments. Brian Switek covered this quite well in his post on it, so I will not review that history. There was a big Museum of the Rockies exhibit about it that toured the USA, and other media attention surrounding it, so Horner’s name became attached to the idea as a result. Other such as Lambe and Colinvaux had addressed it before, but their ideas never seemed to gain as much currency as Horner’s did. But this post is not about that.

What this post is about is a consideration of why this is still an issue that the media report on (and scientists publish on; the two are synergistic of course), if most scientists aware of past debates are in good agreement that a T. rex was like most other carnivores and was opportunistic as a switch-hitting scavenger-predator, not a remarkably stupid animal that would turn down a proper meal that was dead/alive. Indeed, the Nature news piece has a juicy quote from Horner that implies (although I do not know if it was edited or if important context is missing) that he has been in favour of the opportunistic predator-scavenger conclusion for some time. Thus, as Switek’s article notes, even the strongest advocates of the obligate scavenger hypothesis(?) have changed their minds; indeed, that 2011 blog post intimates that this had already happened at least 2 years ago.

For many years, nothing has been published in the main peer-reviewed literature that favours that extreme “obligate scavenger” hypothesis. If I am wrong and there is a scientific debate, where are the recent papers (say within the past 5 years) that are strong, respectable arguments in favour of it? I contend that it is a dead issue. And if it is just about the middle ground; i.e. what percent of its time did a T. rex spend hunting vs. scavenging; we have no clue and may never know, and it’s not a very interesting question.

But who then is feeding off of this moribund equine; this defunct tyranno-parrot?

In thinking about my reply to the journalist over the past 2 days, I am reminded again of my general feeling that this is no longer a question of scientific evidence; the important bit in bold font above. Maybe we just like this “hypothesis” or the “controversy”, or maybe we’re lazy and don’t want to have to hunt for real debates in science.

But who are “the people?” I do not feel that The Public should be blamed; they are the people that The Scientists and The Media ostensibly are seeking to inform about what the state of modern knowledge and uncertainty is in science. So when I get asked about the controversy after a public lecture, I always try to go into detail about it. I don’t sigh and say “go Google it”. Nor do I do this to a journalist. Indeed, I’ve generally headed this issue off at the pass and added a blurb to press releases/webpages explaining my T. rex research to explain how it relates to the non-controversy; example here.

I have to begin turning my finger of accusation away from scientists and toward some of the media, because they must play a huge role in the shennanigans. Yes, scientists should know better then to play this up as a valid, heated, modern controversy. That is true. Yet I have a feeling that the balance of blame should also fall heavily on the side of media (general and science news) that continue to report on this issue uncritically as a real controversy. Thus the general public thinks it still is, and scientists/journals keep issuing papers/press releases that it is, leading to more reporting on this “controversy”, and the beast refuses to die. Switek’s article is a good counter-example of balanced coverage with clear application of critical thinking.

This is trivially different from other non-controversies in palaeontology such as whether birds evolved from a subgroup of theropod dinosaurs and hence are dinosaurs by virtue of descent (consensus = yes). So it is reflective of a broader problem of not calling a spade a spade.

And it’s embarassing, to a scientist, as my quote above expressed, to see dead controversies trotted out again and again, feeding the public perception that they are not dead.

That’s what leaves me frustrated. When do the shennanigans end?

I am reminded of a quote from a Seinfeld episode:

“Breaking up is like knocking over a Coke machine. You can’t do it in one push. You gotta rock it back and forth a few times, and then it goes over.”- Jerry, from the episode “The Voice”.

But this predator/scavenger relationship-from-hell leaves me, as a specialist working in this general area, feeling like I am trapped under that fridge. Help!

That’s why I started off this long post talking about feeling deflated, or disappointed, when asked this question. I do feel that way. I have to admit, I sometimes evenfeel that way when a sweet young kid asks me that question. Deep inside, I wish they wondered about something else. I wish that science had reached them with a deeper, more contemporary question. But when a journalist asks me how I feel about a new paper that revisits the “controversy”, I feel embarassed for palaeontology. Can’t we get past this? It makes us look so petty, mired in trivial questions for decades. But we’re not like that. This is a dynamic, exciting, modern field, but every news story about non-issues in palaeontology just perpetuate bad elements of palaeontology’s image.

To the scientists– why don’t we put our foot down more and say enough is enough, this is a dead issue? We have a role not only in peer review, but also in communicating our views about published work to the media when asked (AND when not asked, as in this blog post). But if you call them on it, do they listen? Which brings me to…

To the media (science/general journalists etc; I know this is a huge category and please don’t think I am blaming 100% of journalists or assuming they are all the same; they are not!)– if scientists tell you that a “controversy” is not such, at what point do you accept their judgement and kill the story, or at least use that quote? Does that ever happen? In what way are you at the mercy of senior editors/others in such issues? What power do you have? Is a shift in the balance of editorial power needed, or even achievable, in your case or in good exemplar cases? I’d really like to hear your experiences/thoughts. I am sure there is a lot I am not understanding, and I know many journalists are in a tough situation.

To the public– You’re often being misinformed; you are the losers in this issue. How do you feel about all it? (While this post focuses on a very tiny issue, the T. rex scavenger/predator unending drama, it is also about a broader issue of how the media perpetuates controversies in science after they have already gone extinct.)

What did this post have to do with freezers? Nothing. I’m just (H)ornery. Although I was once filmed for a planned Discovery Channel film about scientists who find a frozen tyrannosaur in polar regions and have to decide what to do with it before it slips into a chasm and is lost forever. Probably better that this never aired; it was cancelled. Segue to this post.

The Berkeley cast of the Wankel (MOR555) specimen of T. rex. Will we ever see the end of the predator/scavenger non-issue?